Jitka Klimešová, Jiří Danihelka, Jindřich Chrtek, Francesco de Bello and Tomáš
Herben: CLO-PLA: a database of clonal and bud-bank traits of the Central European
flora. Ecology 98: 1179.
INTRODUCTION
One of the defining feature of plants is the modular construction of their bodies. Individual
plant species differ in the type (e.g. root- vs. stem-derived) and degree of this modularity, and
study of these differences is a big challenge for plant ecology. In contrast to the unitary
construction of annual plants and many trees, perennial herbs are usually composed of
multiple shoots that are connected below ground. The connections between these shoots may
disintegrate, causing individual shoots to become independent rooting units and thus resulting
in clonal reproduction (Harper 1977, Jackson et al. 1985, Klimeš et al. 1997). Clonal
reproduction thus provides an alternative pathway to sexual reproduction, ensuring long-term
population persistence, and constitutes one of the most prominent axes of variation in lifehistory traits of perennial herbs (Mogie and Hutchings 1990, Eriksson and Jerling 1990, Grace
1993, Aarssen 2008). Clonal growth plays a key role both in short-distance dispersal and in
persistence within habitats, while also contributing to long-distance dispersal if clonal
offspring become loose and get transported by wind or water. Consequently, plants with a
capacity for clonal growth exhibit characteristic spatial and temporal patterns (Klimeš et al.
1997) with potentially profound impacts on plant population dynamics (Eriksson and
Jakobsson 1998, Winkler et al. 2010).
Clonal growth and reproduction is characterized by a rich variety of plant structures
forming a complex set of plant functional traits that are important for species fitness (Klimeš
et al. 1997, Pan and Price 2001). However, the functional trait data currently used in speciesbased or community-based analyses have only rarely included traits that represent the
potential for clonal growth and multiplication. As this ability is a key attribute of many
perennial herbaceous plants, with consequences for ecosystem functions (Bardgett et al.
2015), the trait data in current use lack an important aspect of the plant life history. This is
due to a number of reasons. In particular, clonal growth is not easily captured by only one
parameter. Different plant groups possess very different organs of clonal growth, which
makes their classification a complicated task. In many plant species, clonal growth is, to an
important degree, environmentally dependent (Sammul 2011), which makes it difficult to
define species-specific information. Finally, organs of clonal growth are often positioned
belowground, which makes the collection of large datasets on them fairly laborious.
Still, the identification of a set of easily measurable traits that describe clonal growth
should be essential for discerning its effects on species distribution and abundance, on
community assembly, and on their consequences for ecosystem functioning. Data on clonality
of individual species can be analysed in terms of their relationships to other traits and
potentially serve to identify broader plant functional types (Herben et al. 2016). Clonality can
play a role in community assembly (Dalgleish and Hartnett 2009, Sosnová et al. 2010, Gough
et al. 2012, Klimešová and Herben 2015). Existing comparative data (Hess 1909, Söyrinki
1938, Gustafsson 1948; Hartmann 1957, Leaky 1981, Klimeš and Klimešová 1999, Tamm et
al. 2001, Klimešová and de Bello 2009) have already yielded a number of important insights,
although in most cases they are limited to a small number of species and/or habitats. In this
paper we are building on earlier attempts to quantify clonal growth traits for larger sets of
species (Klimeš and Klimešová 1999, Klimešová and Klimeš 2008, Klimešová and de Bello
2009) and provide a synthetic dataset characterizing each species in terms of key traits of its
clonal growth. In particular, we use the classification of clonal growth types developed by
Klimeš et al. (1997) and Klimešová and de Bello (2009). However, the earlier datasets
suffered from a number of drawbacks. Namely, they provided rather raw data on individual
species that needed expert evaluation to be used in phylogenetic or community analyses and,
often, required dedicated data manipulation. In contrast to these earlier efforts, we therefore
aim to provide fully interpreted species-level data on individual traits where different records
and data on individual species are synthesized. This provides a dataset in which each species
is characterized by the ecologically most relevant type of its clonal growth and typical
lifespan. The present effort covers a great variety (~2900 species) of temperate species from
Central Europe. Details on the species and the species data are provided below (Metadata,
II.B). Last not but least, we believe that assembling and interpreting such data will show that
it is possible to establish trait databases fully including clonal-related traits, which may serve
as an incentive to collect such data in other regions, which will help to close this gap in our
knowledge of plant life histories.
METADATA
CLASS I. DATASET DESCRIPTORS
A. Dataset identity:
Title: CLO-PLA: a database of clonal and bud-bank traits of Central European flora
B. Dataset identification code:
Suggested dataset identification code: CLO-PLA 3.4
C. Dataset description
1. Principal Investigators:
Jitka Klimešová, Institute of Botany, Institute of Botany, Czech Academy of Sciences, CZ379 82 Třeboň, Czech Republic, [email protected]
Jiří Danihelka, Department of Botany and Zoology, Faculty of Science, Masaryk University,
[email protected]
Jindřich Chrtek, Institute of Botany, Czech Academy of Sciences, CZ-252 43 Průhonice,
Czech Republic
Francesco de Bello, Institute of Botany, Czech Academy of Sciences, CZ-379 82 Třeboň,
Czech Republic and Department of Botany, Faculty of Science, University of South Bohemia,
České Budějovice
Tomáš Herben, Institute of Botany, Czech Academy of Sciences, CZ-252 43 Průhonice,
Czech Republic and Department of Botany, Faculty of Science, Charles University, Benátská
2, CZ-128 01 Praha 2, Czech Republic, [email protected]
Abstract: This dataset presents comprehensive and easy-to-use information on 29 functional
traits of clonal growth, bud banks and lifespan of members of the Central European flora. The
source data were compiled from a number of published sources (see the reference file) and the
authors’ own observations or studies. In total, 2909 species are included (2745 herbs and 164
woody species), out of which 1532 (i.e. 52.7% of total) are classified as possessing clonal
growth organs (1480, i.e. 53.9%, if woody plants are excluded). This provides a unique, and
largely unexplored, set of traits of clonal growth that can be used in studies on comparative
plant ecology, plant evolution, community assembly and ecosystem functioning across the
large flora of Central Europe. It can be directly imported into a number of programmes and
packages that perform trait-based and phylogenetic analyses aimed to answer a variety of
open and pressing ecological questions.
D. Key words: clonal traits; bud-bank traits; Central European flora; Czech flora; functional
ecology; lifespan; vascular plants.
CLASS II. RESEARCH ORIGIN DESCRIPTORS
A. Overall project description
Identity: Traits of clonal growth and bud bank for functional analyses based on lists of plant
species
Originator: Jitka Klimešová
Period of Study: 1994 - 2016
Objectives: To assemble data on traits of clonal growth and bud banks of species in the
Central European flora. We assembled and interpreted data on 29 traits of clonal growth from
published sources and our own field data.
Abstract: This dataset presents comprehensive and easy-to-use information on 29 functional
traits of clonal growth, bud banks and lifespan of members of the Central European flora. The
source data were compiled from a number of published sources (see the reference file) and the
authors’ own observations or studies. In total, 2909 species are included (2745 herbs and 164
woody species), out of which 1532 (i.e. 52.7% of total) are classified as possessing clonal
growth organs (1480, i.e. 53.9%, if woody plants are excluded). This provides a unique, and
largely unexplored, set of traits of clonal growth that can be used in studies on comparative
plant ecology, plant evolution, community assembly and ecosystem functioning across the
large flora of Central Europe. It can be directly imported into a number of programmes and
packages that perform trait-based and phylogenetic analyses aimed to answer a variety of
open and pressing ecological questions.
Sources of funding: Czech Science Foundation (Centre of Excellence PLADIAS, 1436079G), institutional funding by the Institute of Botany.
B. Specific subproject description
1. Species included
The dataset includes a major part of the Central European flora. We included all vascular
plant species regularly occurring in the Czech Republic following Danihelka et al. (2012) and
a number of additional species from the BiolFlor database (Klotz et al. 2002, Kühn et al.
2004). In total, the database encompasses 2909 plant species. Although the primary focus is
on clonal traits, plant species were included regardless of their growth form. The database
thus covers both clonal and non-clonal species. This makes it possible to perform comparative
studies of all plants irrespective of their ability to spread clonally.
The taxonomy and nomenclature generally follows the The Plant List (2013), Euro+Med
(2006 onwards) and the checklist of vascular plants of the Czech Republic (Danihelka et al.
2012). Whenever the taxonomy and nomenclature in these sources disagreed, we consulted
further sources, such as recent taxonomic studies and the most important local floras, namely
Jäger (2011) and Fischer (2008). We also considered the taxonomic concepts adopted in the
most important national and regional floras and checklists as well as those proposed in recent
monographic studies. The names used constitute a practical compromise without any intention
to provide any reference taxonomy or nomenclature. With a few exceptions, we did not
include subspecies and hybrids. Author names are abbreviated following the conventions of
the International Plant Names Index (IPNI 2004 onwards).
Species aggregates (e.g. Rubus fruticosus agg.) were introduced in taxonomically intricate
groups whenever the species concept (narrow vs. broad) was not clearly stated by the authors
of original data. We also adopted species aggregates used in source publications.
2. The traits
The database contains the following broadly defined groups of traits: life history, whole-plant
traits, bud bank traits, traits of clonal growth and root sprouting traits. Within each of these
groups, several specific functional traits are considered. As in all existing functional trait
databases worldwide, not all traits are available for all species. This has two reasons: First,
some traits are defined only for plant species with specific life histories. Traits of (stemderived) clonal growth are defined only for clonal perennial herbs (i.e. not for annuals,
monocarpic perennials, perennial non-clonal and woody plants; the definition of the term
‘woody plant’ is provided below). Whole-plant traits (shoot cyclicity and primary root
presence) are defined for all herbaceous plants (excluding woody plants). Bud bank traits and
the presence of root sprouting are defined for all plants. The other two traits of root sprouting
(Role and Position) are defined only for root-sprouting species. Bud bank depth is defined
only for species with non-zero bud bank. Second, we do not possess complete information on
all traits in some species, due to the quality of source data or absence of personal
observations. These are the missing values that do not fall into the categories listed above in
this paragraph. They typically constitute only a small fraction of meaningful values: less than
1% for life-history, bud-bank and root-sprouting traits, less than 4% for traits of clonal
growth, 7% for shoot lifespan (cyclicity), and 1% for primary root presence. The only trait
with a larger proportion of unknown values is branching (34%).
The source data on the reported traits were obtained from various existing sources (see
the list of references in the CLO-PLA-reflist.txt file), by the interpretation of
descriptions and drawings in floras and research articles, and based on the first author's
personal observations. These data are stored in the CLO-PLA3 database (Klimešová and
Klimeš 2006, 2008), on which the current synthesis is based. In the CLO-PLA3 database, in
fact, multiple sources for most species are available, but for the present synthesis they have
been carefully screened and summarized. The method of synthesizing these data is described
separately for each trait or group of traits below. With the exception of life history, the basic
unit for the assessment of each individual trait was the shoot (product of the apical meristem
with a potential to flower).
2.1. Life history
Life-span/ life-history data were based on the BiolFlor database and corrected according to
data in the CLO-PLA database and in Jäger and Werner (2002). If two possibilities were
reported, we selected the shorter lifespan. Small shrubs (Chamaephytes according to
Raunkiaer) were not included among woody plants, as they are functionally closer to
perennial herbs than to shrubs and trees.
2.2. Whole-plant traits
Branching type of stem-derived organs of clonal growth. The branching type of perennial
herbs determines whether individuals possess two different shoot types (flowering and sterile)
or only one shoot type (which can potentially flower). In sympodially branched plants, all
shoots are identical in their construction, replace each other during the ontogeny of the
individual, and all of them can potentially flower. By contrast, monopodially branched plants
possess two shoot types, one of which never flowers whereas flowering shoots arise from
axillary buds of the non-flowering shoot. Finally, ferns and lycopodioids can possess
dichotomous branching, which is functionally similar to monopodial branching. The data
reported here are based on individual observations in the CLO-PLA3 database; if more types
are reported for one species, the most frequently observed type is given here.
Shoot life span (cyclicity). This trait determines the number of years from the emergence of
the aboveground part of the shoot to flowering and fruiting (see Serebryakov 1952,
Klimešová et al. 2016). We distinguish monocyclic plants with shoots living for one season
only, and di- and polycyclic plants with shoots that typically live longer than one year and
never flower in the first year. This has a number of demographic consequences: Monocyclic
plants typically do not possess a leaf rosette and all aboveground shoots in a population are of
the same age. By contrast, di- and polycyclic plants possess a basal leaf rosette and their shoot
populations include flowering and sterile shoots of several cohorts at the same time. Shoot
lifespan (cyclicity) has a different meaning depending on the type of branching (see above). In
sympodially branched plants, cyclicity refers to all shoots; in monopodially branched plants, it
refers only to flowering shoots. The data are based on individual observations in the CLOPLA3 database. If more types are reported for one species, the most frequently observed type
is given here. Furthermore, di- and polycyclic plants are merged into a single type (denoted
polycyclic), as analysis based on morphological traits permits the identification of shoots with
a cyclicity of one year (monocyclic) from those that live longer (di- and polycyclic).
Primary root presence. If the primary root is the only root for the whole life of a plant, the
plant is unable to form roots on stems (i.e. adventitious roots) and is therefore non-clonal
(unless it is able to form adventitious buds on roots). If the primary root is replaced during
ontogeny by adventitious roots formed on belowground parts of the stem, the plant is able to
grow clonally. In some species the primary root can split in later life stages, also yielding
several independent individuals. Some species form adventitious roots only under specific
conditions (soil moisture, root injury, or old age). The data reported here are based on
individual observations in the CLO-PLA3 database; if more types are reported for one
species, the most frequently observed type is given here.
2.3. Bud-bank traits
The term ‘bud bank’ denotes all inactive (dormant) buds on the plant body that can give rise
to new shoots (Klimešová and Klimeš 2007). The most important part of the bud bank for
plants in temperate climates are buds located out of the reach of frost or drought (Raunkiaer
1934), i.e. on the soil surface or below ground. We therefore report only data on buds located
at or below the soil surface.
The number of buds on plant organs located at different soil depths was assessed according to
a simple method based on morphological characters (Klimešová and Klimeš 2007). The
assessment was based on the assumption that each leaf (or leaf scale) axil contains a bud. The
assessment of bud numbers was done in categories (0, 0–10, >10 buds per shoot; Klimešová
and Klimeš 2008), which were represented for further calculations by the centers of these
categories (0, 5, 15 buds per shoot). The final value was calculated as the mean of all these
values for the given species and given soil depth. In addition, some plants possess the ability
to form adventitious buds on the root or hypocotyl. As root buds cannot be counted (they are
formed freely along the root), for categories that include root buds, 15 buds were arbitrarily
added per each 10 cm of depth as a practical compromise based on personal observations of
the authors (see also Klimešová and Klimeš 2007). We use the term ‘root buds’ as a general
term denoting both buds on roots and the hypocotyl.
2.4. Traits of clonal growth
Clonal growth is defined as growth of the plant body that can potentially lead to the formation
of physically independent asexual offspring. A morphological prerequisite for such growth is
the formation of adventitious roots on stems and/or adventitious buds on roots (Groff and
Kaplan 1988). This general process takes place at clonal growth organs with different degrees
of specialization, and some species can even possess several independent types of such
organs. With the exception of the type of clonal growth organs (CGO, see below), all traits
reported in this section are reported only for herbs that possess the ability to spread clonally.
The type of clonal growth organs is reported for all species (including woody species) that are
able to spread clonally.
Type of clonal growth organs (CGO). While some species can possess several independent
types of clonal growth organs, the present data report only one, selected as being the most
important for the life cycle of the species, that is the one that produces the greatest number of
offspring and/or enables the individual to spread its offspring over larger distances. All clonal
traits are then reported for this organ type.
CGO types are morphological categories that are defined based on several parameters
(Klimešová and Klimeš 2006, 2008; Table 1): (i) bud bearing organ (BBO), i.e. which organ
bears buds that give rise to the given CGO (shoot or root); (ii) position, i.e. how the organ is
positioned relative to the soil surface; and (iii) storage, i.e. the organ where resources are
stored (shoot, root, leaf).
Table 1. Classification of types of organs of clonal growth (CGO).
CGO type
Numeric coding
in the database
Bud bearing
organ
Position
Storage
Horizontal stem
1
shoot
aboveground
shoot
Turions
2
shoot
aboveground
(submerged)
bud
Stem fragments
5
shoot
aboveground
(submerged)
shoot
Budding
6
shoot
aboveground
(submerged)
plantlet
Epigeogenous
rhizome
9
shoot
initially
aboveground, later
belowground
shoot
Hypogeogenous
rhizome
10
shoot
belowground
shoot
Stem tuber
12
shoot
belowground
part of the shoot
Bulb
13
shoot
belowground
leaf
Root sprouting
15
root
belowground
root
Root tuber
16
shoot
belowground
root
Persistence of clonal growth organs. The persistence of a clonal growth organ determines the
lifespan of the physical connection between mother and daughter shoots. Because
morphological analysis does not permit the identification of such lifespan beyond a few years
only, the reported values are based on truncated values. The persistence of the connection was
assessed in categories (0.5, 1.5 and 4 years); the reported values are means of all records for
the given species. As a result, the data are effectively truncated by the value of 4 years, which
should be interpreted as 4 years or more. Analyses using this trait should therefore take this
truncation into account.
Number of clonal offspring. Number of offspring shoots produced per parent shoot per year.
The numbers of clonal offspring were estimated within categories (<1, 1, 2-10, 10+), which
were further represented by mean values of their ranges (0.5, 1, 6, and 15). The reported
values are means of these value in all records for the given species (Klimešová and Klimeš
2006).
Lateral spreading distance. Distance between parental and offspring shoots. Lateral spreading
distances were estimated within categories (<1 cm, 1-25 cm, 25+ cm), which were further
represented by mean values of their ranges (0.5 cm, 13 cm, 50 cm). The reported values are
means of these values in all records for the given species (Klimešová and Klimeš 2006).
Clonal index. An aggregate measure of the ability of a species to spread clonally (Johansson
et al. 2011). It is based on categories of traits denoted as Lateral spreading distance (<1 cm,
1-25 cm, 25+ cm) and Number of clonal offspring (<1, 1, 2-10, 10+). These categories were
assigned ordinal values (1, 2, 3, 4). Plants with small and dispersible offspring were assigned
the lateral spread score of 4. The clonal index is defined as the sum of ordinal values of both
traits. The data reported here are based on individual observations in the CLO-PLA3
database; if more categories of either the number of offspring or the lateral spreading distance
were reported for a species, the most frequently observed type is given here (Klimešová and
Klimeš 2006).
Presence of freely dispersible organs of clonal growth. Another form of clonality is the
formation of freely dispersible clonal offspring, i.e. new individuals that are separated from
the mother shoots very shortly after their formation and before they develop roots attaching
them to the soil. They are dispersed by wind, water, animals or other agents. Typical
examples are fragmenting shoots, bulbils, turions or fragments of water plants. The data
reported here are based on individual observations in the CLO-PLA3 database (Klimešová
and Klimeš 2006).
2.5. Traits of root sprouting
The ability to form adventitious buds on roots is an alternative mode of bud bank formation
and clonal growth. A plant is marked as a root sprouter if there is at least one confirmed
positive report.
Position of root buds. In contrast to shoot buds, which are always located in a leaf axil,
adventitious buds on roots (or the hypocotyl) are not located at any specific positions; they
occur only on some root types or may appear only after the individual is injured. We list the
hypocotyl only if the roots are found exclusively on it; we report the main root if the buds are
never formed on lateral roots, and we report lateral roots if root buds are potentially found on
all three types of bud-bearing organs.
Role of root buds in the plant life history. Root buds have different roles in individual species.
In some species, they occur only after the plant is injured and play a role in post-injury
regeneration (regenerative role). In some other species, these buds are formed without such
external stimulus and thereby increase the number of shoots and thus the number of offspring,
both clonally and by seed, but are not necessary for completing the life cycle; we denote them
as facultative. In a few species, the formation of root buds is necessary for the completion of
their life cycle. They develop in all individuals and condition their survival/flowering or
overwintering (Klimešová 2007). We denote them as obligate root sprouters. This ability is
hierarchical: Obligate root sprouters are able to resprout also as facultative or regenerative
root sprouters, and facultative root sprouters are able to regenerate after injury with the help
of adventitious buds.
CLASS III. DATASET STATUS AND ACCESSIBILITY
A. Status
Latest update: September 2016
Latest Archive date: September 2016
Metadata status: Metadata are complete and are stored with the data.
Data verification: Data were checked during entry.
B. Accessibility
Storage location and medium: All data and metadata are stored with this publication in
Ecology. Further, all digital data and metadata are stored on the principal investigators’
computers and on computer hard drives at the Institute of Botany, Czech Academy of
Sciences, Průhonice and Třeboň.
Contact persons:
Jitka Klimešová, Institute of Botany, Institute of Botany, Czech Academy of Sciences, CZ379 82 Třeboň, Czech Republic, [email protected]
Tomáš Herben, Institute of Botany, Czech Academy of Sciences, CZ-252 43 Průhonice,
Czech Republic and Department of Botany, Faculty of Science, Charles University, Benátská
2, CZ-128 01 Praha 2, Czech Republic, [email protected]
Copyright restrictions: None, the dataset is freely available for scientific use.
Proprietary restrictions: None. The data originator would appreciate notification when and
how the data are used. When used in published analyses, this paper should be referred to as
the data source.
a. Release date: To be specified by the time of publication
b. Citation: This paper
CLASS IV. DATA STRUCTURAL DESCRIPTORS
A. Dataset file: The dataset is downloadable as a single archive, DataS1.zip. This file
contains three data files described below.
IV.A.1 Data file name: CLO-PLA-traits.txt
This file is a tab-delimited ASCII file containing the list of species with trait data.
Header information:
Abbreviation
Full name
Trait
type1)
Definition
woody
Woody plant
Cat
1 - woody plants, excluding small shrubs (chamaephytes) and
suffruticose plants, 0 - otherwise
annual
Annual plant
Cat
1 - plants typically living for one season. Includes winter annuals, 0
- otherwise.
Nonclonal
perennials
Cat
1 - plants that do not multiply vegetatively (typically possessing a
primary root) and are neither annual (i.e. live for more than one
year) nor woody ("nonclonal plants"), 0 - otherwise
monocarpic
Monocarpic plants
Cat
1 - nonclonal perennials that flower only once in their lifetime, 0 otherwise.
polycarpic
Polycarpic plants
Cat
1 - nonclonal perennials that flower several times in their lifetime, 0
- otherwise.
Clonal plants
Cat
1 - plants that live longer than one year, multiply vegetatively and
flower several times in their lifetime, 0 - otherwise
branching
Branching type
Cat
monop – monopodial branching, symp – sympodial branching, dich
– dichotomous branching (functionally monopodial). Indicated only
for polycarpic (clonal and non-clonal) plants.
cyclicity
Shoot cyclicity
Cat
1 - monocyclicity prevailing, 2 - otherwise (dicyclic or polycyclic
shoots prevailing).
Primary root
present
Cat
1 - primary root is present, 0 - otherwise
BB0
Bud bank at soil
surface - number
Cont
Number of stem-derived buds at the soil surface.
BB0_mn10
Bud bank in soil
depth 0-10 number
Cont
Number of stem-derived buds in the soil in the depth range of 0-10
cm.
BB_gtmn10
Bud bank in soil
depth below 10 number
Cont
Number of stem-derived buds in the soil below 10 cm.
Bud bank at soil
surface (all buds) number
Cont
Number of all buds (stem-derived and root-derived) at the soil
surface.
Life history
perennialnonclon
al
clonal
Whole-plant
traits
Primaryroot
Bud-bank traits
BB0R
BB0_mn10R
Bud bank in soil
depth 0-10 (all
buds) - number
Cont
Number of all buds (stem-derived and root-derived) in the depth
range of 0-10 cm.
BB_gtmn10R
Bud bank in soil
depth below 10 (all
buds) - number
Cont
Number of all buds (stem-derived and root-derived) below 10 cm.
BBsize
Total stem-derived
bud bank size
Cont
Total number of buds, calculated as
BB0 + BB0_mn10 + BB_gtmn10
BBdepth
Mean stem-derived
bud bank depth
[cm]
Cont
Mean bud bank depth calculated as
(BB0 + BB0_mn10 *5 + BB_gtmn10 *15) /
(BB0 + BB0_mn10 + BB_gtmn10)
BBRsize
Total bud bank size
(including root
buds)
Cont
Total number of buds, calculated as
BB0R + BB0_mn10R + BB_gtmn10R
Mean bud bank
depth (including
root buds) [cm]
Cont
Mean bud bank depth calculated as
(BB0R + BB0_mn10R *5 + BB_gtmn10R *15) /
(BB0R + BB0_mn10R + BB_gtmn10R)
Cat
Morphological type of clonal growth organ (for the coding see
Table 1). If a species possesses more than one clonal organ type, the
dominant one (i.e. playing most important ecological role) is given.
Persistence of
connections [yrs]
Cont
Persistence of clonal connections in years.
Number of
offspring
Cont
Number of offspring shoots per parent shoot.
offspring_wsmall
Number of
offspring including
small offspring
Cont
Number of offspring per parent shoot including offspring of small
size. Small offspring are defined as clonal offspring for which it
took more years to attain a size comparable with other clonal
offspring of the plant; they usually resemble seedlings.
spread
Lateral spread [cm]
Cont
Lateral spreading distance of clonal growth organs. Values are based
primarily on offpring of normal size, but were edited when
necessary based on an educated guess using also small offspring
(freely dispersible clonally formed organs are excluded).
clonalindex
Clonal index
Ord
Sum of ordinal values of categories of Lateral spread and Number of
clonal offspring
dispersibility
Freely dispersible
diaspores present
Cat
Presence of freely dispersible clonal organs. 1 – present, 0 – absent.
Rsprouter
Existence of the
root bud bank
Cat
1 – root bud bank reported for the species; 0 – no reports of root bud
bank or root bud bank absent
PositionRB
Root bud bank
position
Cat
1 - hypocotyl, 2 - main root, 3 - lateral roots
Root bud bank role
Cat
F - facultative, R - regenerative, O - obligate
BBRdepth
Traits of clonal
growth
finalCGO
persistence
offspring
Traits of root
sprouting
RoleRB
1)
Cat – categorical, Cont – continuous, Ord - ordinal
IV.A.2 Data file name: CLO-PLA-refs.txt
This file is a tab-delimited ASCII file containing the list of species (in the same order as in the
CLO-PLA-traits.txt file) with references used to obtain information on individual
species.
Column header
Definition
Species name, identical to that in the file CLO-PLA-
Species_name
traits.txt
References
Data sources for the species
IV.A.3 Data file name: CLO-PLA-reflist.rtf
This file is an ASCII file containing the list of references referred to in the file CLO-PLArefs.txt.
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